SCIENCE CHINA Earth Sciences • RESEARCH PAPER • December 2010 Vol.53 No.12: 1836–1848 doi: 10.1007/s11430-010-4089-6 Cranial morphology of the Silurian sarcopterygian Guiyu oneiros (Gnathostomata: Osteichthyes) QIAO Tuo1,2 & ZHU Min1* 1 Key Laboratory of Evolutionary Systematics of Vertebrates, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China; 2 Graduate School of Chinese Academy of Sciences, Beijing 100049, China Received April 6, 2010; accepted July 13, 2010 Cranial morphological features of the stem-group sarcopterygian Guiyu oneiros Zhu et al., 2009 provided here include the dermal bone pattern and anatomical details of the ethmosphenoid. Based on those features, we restored, for the first time, the skull roof bone pattern in the Guiyu clade that comprises Psarolepis and Achoania. Comparisons with Onychodus, Achoania, coelacanths, and actinopterygians show that the posterior nostril enclosed by the preorbital or the preorbital process is shared by actinopterygians and sarcopterygians, and the lachrymals in sarcopterygians and actinopterygians are not homologous. The endocranium closely resembles that of Psarolepis, Achoania and Onychodus; however, the attachment area of the vomer pos- sesses irregular ridges and grooves as in Youngolepis and Diabolepis. The orbito-nasal canal is positioned mesial to the nasal capsule as in Youngolepis and porolepiforms. The position of the hypophysial canal at the same level or slightly anterior to the ethmoid articulation represents a synapmorphy of the Guiyu clade. The large attachment area of the basicranial muscle indi- cates the presence of a well-developed intracranial joint in Guiyu. Sarcopterygii, Osteichthyes, Cranial morphology, homology, Silurian, China Citation: Qiao T, Zhu M. Cranial morphology of the Silurian sarcopterygian Guiyu oneiros (Gnathostomata: Osteichthyes). Sci China Earth Sci, 2010, 53: 1836–1848, doi: 10.1007/s11430-010-4089-6 The primitive fish Guiyu oneiros [1] from the Silurian of nopterygians and sarcopterygians, and between osteichthy- China represents the oldest articulated osteichthyan, and ans and non-osteichthyan gnathostomes, thus forcing a re- provides unique insights into the character transformation appraisal of the sequence of character acquisition in the that resulted in the standard set of sarcopterygian features early evolution of gnathostomes [15–17]. However, the [2]. Prior to the discovery of Guiyu, the earliest fossil record early diversification of gnathostomes remains unclear due to of the osteichthyans was represented only by isolated frag- the incompleteness of the fossil record and the lack of the mentary remains, such as isolated scales and teeth [3–10]. detailed anatomical studies on some known taxa. The articulated material of Guiyu reinforces novel interpre- The earlier report of Guiyu oneiros [1] only provided a tations of basal sarcopterygian morphology previously brief description of the taxon with emphasis on the mosaic based on disarticulated specimens of Psarolepis [11]. The of primitive gnathostome characters, and placed Guiyu, mosaic of characters in some early fishes [11–14] has sig- Psarolepis, and Achoania into a monophynetic group, the nificantly bridged the morphologicial gaps between acti- Guiyu clade. Here we offer a detailed description of the cranial morphology based on some newly-prepared disar- ticulated specimens assigned to Guiyu oneiros, as well as *Corresponding author (email: [email protected]) those illustrated in ref. [1], and provide a restoration of the © Science China Press and Springer-Verlag Berlin Heidelberg 2010 earth.scichina.com www.springerlink.com QIAO Tuo, et al. Sci China Earth Sci December (2010) Vol.53 No.12 1837 skull roof bone pattern in the Guiyu clade [1]. All speci- ing of the posterior cranial portion. In the rest of early sar- mens were recovered from the muddy limestone of the copterygians, the anterior cranial portion is usually much Kuanti Formation (Late Ludlow, Silurian) in Qujing, Yun- longer than the posterior cranial portion as in Psarolepis. nan, China [1]. In light of new anatomical information, the Considering the state in Meemannia [23], which is immedi- homology is examined of certain cranial bones including the ately basal to the Guiyu clade, and early actinopterygians preorbital of basal actinopterygians and the lachrymal of [24], we suggest that the equal length of the two cranial sarcopterygians. The present paper follows the terminology portions in Guiyu might represent a derived feature. of Westoll [18], only to align with current majority usage. The parietal shield is square shaped and has its maximum width at the level of the postorbital corners. The postparietal shield is broad posteriorly and is broadest at the level of the 1 Cephalic exoskeleton suture between the supratemporal (St, Figures 1, 2(f) and 1.1 General features of skull roof 3(f)) and tabular (Ta, Figures 1, 2(f) and 3(f)), and then narrows slightly. Its posterior margin is straight, with an The holotype (IVPP V15541) has a skull roof of about 4.5 inconspicuous medial projection. cm in length, which is divided into the parietal and postpa- The dermal bones on the skull roof of Guiyu contain rela- rietal shields by the dermal intracranial joint (Figure 1(a)). tively few discrete elements, which may represent the The parietal shield of the holotype looks broader and shorter primitive sarcopterygian condition. They are ornamented than those disarticulated ones (Figures 2 and 3), whose with vermiculate ganoine ridges (Figures 1, 2, 4(a), 4(b) and length ranges from 1.9 to 2.0 cm. As evidenced by the ar- 5(a)), which for the most part have fused into an irregular ticulated specimen, the length ratio between the anterior and maze-like configuration. Ganoine comprises a glossy outer posterior cranial portions in Guiyu is about 1.0, which is dermal coating of single or multi-layered enamel. It was smaller than that in Psarolepis (1.6–1.7) [19]. In onycho- once considered a characteristic feature of basal actinop- donts [20, 21] and derived porolepiforms such as Glyptole- terygians [14, 24, 26–29]. Richter and Smith [30] noted the pis and Holoptychius [22], the length ratio between two presence of ganoine-like tissue in acanthodians and basal portions is smaller than 1.0, evidently due to the lengthen- sarcopterygians, suggesting that ganoine might be plesio- Figure 1 Guiyu oneiros Zhu et al. 2009. (a) Close-up view of the head in dorsal view, holotype, V15541; (b) Reconstruction of the skull roof based on all available data. It, intertemporal; ano, anterior nostril; Dsph, dermosphenotic; Et, extratemporal; Eta, accessory extratemporal; ioc, infraorbital canal; lc, main lateral-line canal; lR, lateral rostral; mpl, middle pit-line; mR, median rostral; Na1, nasal 1; Na2, nasal 2; Pmx, premaxillary; Pa, parietal; pi, pineal foramen; pno, posterior nostril; Ppa, postparital; ppl, posterior pit-line; Pr, postrostral; Pro, preorbital; So, supraorbital; soc, supraorbital canal; St, supratemporal; Ta, tabular. 1838 QIAO Tuo, et al. Sci China Earth Sci December (2010) Vol.53 No.12 morphic for crown osteichthyans. Schultze [8] indicated the 2(a)–(e), 3(a)–(e), 4(b) and 5), excluding it from the occlu- differences in surface structure between the typical ganoine sal margin of the jaw, as in early actinopterygians [24], in actinopterygians (with surface elevations or microtuber- Youngolepis [32], Powichthys [33, 34], and other crown cles) and the enamel of cosmine in sarcopterygians (with sarcopterygians. The premaxillary is ribbon-shaped. It depressed hexagons marking the borders of adjacent cells); gradually increases in height posterolaterally until it meets however, this distinction is not absolute since the ganoine the suture between the lateral rostral and the preorbital (Pro, without microtuberlces is present in some actinopterygians, Figures 1(b), 3(a)–(e), 4(b) and 5(b)), and then narrows be- such as Cheirolepis trailli [30]. Whether the ganoine sur- neath the preorbital. This marks an additional departure face of Guiyu oneiros or the enamel surface of the dermal from Psarolepis, in which the premaxillary keeps increasing skeleton in other stem sarcopterygians [11, 13, 19, 23, 31] in height posterolaterally and forms a preorbital process bears the microtubercles or not needs further investigation. (pr.Pro, Figure 4(d)) to enclose the posterior nostril (pno, Figures 1, 2(a)–(e), 3(a)–(e), 4(b) and 5). As in Psarolepis 1.2 Parietal shield and Onychodus, the premaxillary bears two types of teeth. The ventral portion bears a row of large teeth at the mouth The bone pattern is hardly discernible in the holotype (Fig- margin. These conical teeth are slender than those in ure 1(a)); however, some disarticulated skull specimens Psarolepis and Achoania, and exhibit the regional variation. (Figures 2 and 3) show distinct bone sutures or patterned In cross section, the teeth on the posterolateral portion of ridges around ossification center, based on which we outline the premaxillary (t.Pmx, Figure 6(d)) display a narrow el- the bone pattern of the parietal shield (Figure 1(b)). lipse shape at the base, and are longer than those close to The upper jaw, in anterior view with an indistinct median the midline of skull roof. Dorsal to this row of larger teeth projection (Figure 4(a) and 4(b)), exhibits a curvature that and at the margin of the dermal ornamentation of the pre- looks intermediate between the somewhat horizontal con- maxillary is a row of minute teeth. figuration in Achoania (Figure 4(c)) and the M-shaped con- The median rostral (mR, Figures 1(b), 3(a)–(e), 4(b) and figuration in Psarolepis (Figure 4(d)). The premaxillaries 5(b)) is lyrate in shape and occupies about 1/4 of the length (Pmx, Figures 4(a), 4(b), 6 and 7(a)–(c)) contact each other of the parietal shield. Anteriorly, it inserts between the pre- at the midline ventral to the median rostral (mR, Figures 1, maxillaries, but does not contribute to the occlusal margin Figure 2 Guiyu oneiros Zhu et al.